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Determinants of Cell Cycle Progression in Human Mammary Epithelial MCF12 Cells

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Determinants of Cell Cycle Progression in Human Mammary Epithelial MCF12 Cells Determinants of cell cycle progression in human mammary epithelial MCF12 cells Alexandra Ouertani Thesis submitted for the degree of Doctor of Philosophy UCL School of Pharmacy University of London 2012 i This thesis describes research conducted in the UCL School of Pharmacy between October 2008 and September 2011 under the supervision of Prof Andreas Kortenkamp and Dr Elisabete Silva. I certify that the research described is original and that any parts of the work that have been conducted by collaboration are clearly indicated. I also certify that I have written all the text herein and have clearly indicated by suitable citation any part of this dissertation that has already appeared in publication. Signature: Date: ii ACKNOWLEDGMENTS First of all, I would like to thank my supervisor Prof Andreas Kortenkamp for giving me the opportunity to undertake this graduate work. Thank you for supporting me in many ways throughout the years, for your time, your positive attitude and the countless discussions we had about this work! I am also very grateful to my second supervisor Dr Elisabete Silva for her constant support, her practical advice and her fresh view on my work. I would like to thank all members of the Centre for Toxicology, who were great colleagues, and all of you have contributed in making my PhD the best time of my life! I wish to thank especially Dr Sibylle Ermler, Dr Richard Evans and Dr Frankie Orton for taking their time to discuss my work, show me new techniques, or to have a much needed break! I would like to thank Dr Erika Rosivatz, Dr Ines da Costa Rocha and Dr Sinikka Rahte. You cheered me up when I doubted myself, and were there for me in many different ways. Thank you for all of this, but mostly for your friendship! I am very grateful to my family, especially my parents and my brother, who were so patient with me during the past years. Finally, I thank my better half Zied, for his encouragement, support and advice during this time. Thank you for always finding the right words to get me through the bad times, and for sharing the good times with me! iii ABSTRACT Cancer of the mammary gland is the most common type of cancer in women worldwide, and the vast majority of breast cancers originate from a cluster of malignant cells in the epithelial tissue of the breast, which initially confines the ductal carcinoma in situ. Research has shown that the signalling pathways that increase differentiation and maintain proliferation in normal epithelial cells are of utmost importance for sustaining this barrier against malignant cells. As a model for normal mammary epithelial cells, the MCF-12A cell line was used to determine factors that are required for cell cycle progression of these cells. A discontinuous treatment assay was developed in which the MCF-12A cells were treated with epidermal growth factor (EGF) and insulin at two distinct times to induce cell cycle re-entry. The use of these chemically defined growth factors enabled us to determine that continuous stimulation with mitogenic factors is not required for these cells to re-enter the cell cycle. An initial activation of the MAP kinase pathway and an up-regulation of the transcription factor c-Myc, followed by activation of the PI3K pathway, resulted in full competence to progress into S phase. The order in which the growth factors were applied, and thus the sequence in which the subsequent proteins were triggered, was of great importance for successful S phase entry. We found that estradiol (E2) was unable to induce the factors necessary for cell cycle progression. Furthermore, we report for the first time that E2 did not affect estrogen- regulated genes which normally are under the control of a ligand-bound estrogen receptor (ER). We suggest that the mechanism by which the ligand-activated ER usually interferes with the estrogen responsive element in the promoter region of the target genes is defective in the MCF-12A cell line. The results presented here may contribute to new approaches in chemotherapy, taking advantage of the diverse molecular mechanism in place for cell cycle progression and proliferation in malignant cells compared to normal mammary epithelial cells. iv TABLE OF CONTENTS Acknowledgments .............................................................................................................. iii Abstract .............................................................................................................................. iv List of Figures................................................................................................................... xiii List of Tables .................................................................................................................... xvi List of Abbreviations ...................................................................................................... xviii CHAPTER 1: INTRODUCTION ................................................................... 1 1 GENERAL INTRODUCTION ...................................................................................... 1 2 CANCER OF THE MAMMARY GLAND ................................................................... 1 2.1 Risk factors for breast cancer......................................................................................... 2 2.2 Signalling between normal and malignant cells for the transition from in situ to invasive carcinoma ...................................................................................................................... 3 3 MAMMARY GLAND EPITHELIUM .......................................................................... 6 3.1 The function of epithelial cells in the mammary gland .................................................. 6 3.2 Signalling for cell cycle progression and proliferation .................................................. 7 4 AIMS ............................................................................................................................ 9 5 THESIS OUTLINE ..................................................................................................... 10 CHAPTER 2: MATERIAL AND METHODS ............................................. 12 1 LIST OF CHEMICALS ............................................................................................... 12 2 ROUTINE CELL CULTURE ...................................................................................... 14 2.1 Routine maintenance of cells ........................................................................................ 14 2.2 Media ............................................................................................................................ 14 2.3 Sub-culturing (passaging) ............................................................................................. 15 2.4 Cryopreservation and resurrection from cryogenic stocks ......................................... 15 2.5 Charcoal-dextran treatment of serum ......................................................................... 16 3 GROWTH FACTORS ................................................................................................. 16 4 INHIBITORS .............................................................................................................. 17 5 FLOW CYTOMETRY ................................................................................................ 17 v 5.1 Discontinuous exposure assay ...................................................................................... 17 5.1.1 Seeding of MCF-12A cells ......................................................................................... 17 5.1.2 Serum depletion for induction of quiescence ............................................................... 18 5.1.3 Incubation with growth factors.................................................................................... 18 5.1.4 Incubation with inhibitors ........................................................................................... 18 5.1.4.1 Inhibitors tested during the first pulse ................................................................... 18 5.1.4.2 Inhibitors tested during the second pulse .............................................................. 19 5.2 Sample preparation for flow cytometric analysis ........................................................ 19 5.3 Staining of cell DNA ..................................................................................................... 19 5.4 Analysis on flow cytometer ........................................................................................... 19 5.5 Data analysis with MACSQuantify™ software ........................................................... 20 5.6 Data analysis with FlowJo software ............................................................................. 20 6 IMMUNOBLOTTING ................................................................................................ 20 6.1 Discontinuous exposure assay ...................................................................................... 20 6.1.1 Seeding of MCF-12A cells ......................................................................................... 20 6.1.2 Serum depletion, incubation with growth factors and inhibitors ................................... 20 6.2 Sample preparation for immunoblotting ..................................................................... 21 6.2.1 Determination of protein concentration and protein separation with SDS-PAGE ......... 21 6.2.2 Transfer .....................................................................................................................
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